Index
Aardwolf, see Proteles cristatusAbsolute configuration, 306–307Acanthomyops (ant), 5Accessory olfactory bulb (AOB),
165, 183, 184, 201(5R,6S)-6-Acetoxy-5-hexadecanolide, 75Acomys cahirinus (spiny mouse)
kin recognition cues, 110Acroneuria carolinensis (stonefly), 236Across-fibre patterning, 171Actias luna (silk moth)
antenna as sieve or paddle, 167Active space, 209
diffusion in air and water, 210–212multi-component pheromones,
212, 214trail, 215
Acyrthosiphon pisum (pea aphid)suicide hypothesis, 152
Adrenal glandsfemale mice, Lee–Boot effect, 195
Aeolidia papillosa (nudibranch sea-slug)predator marked by prey
pheromone, 148Aeshena umbrosa (dragonfly nymph)
predator, 157African mole rats, see Heterocephalus
glaber (naked mole rat)Aggregation pheromones, 74
aposematic insects, 75commercial exploitation
managing bark beetles, 266stopping marine fouling, 266
dilution of risk, 75ecophysiological benefits, 79
assembly pheromone and ticks, 80
overwintering garter snakes, 80intraspecific eavesdropping, 236
males only signal until femalescome, 82
Schistocerca gregaria(desert locust), 76
in space
oviposition pheromones, 75, 76, 77
Panulirus interruptus (lobster), 75settlement of marine
invertebrates, 26, 77–79in time (synchronisation)
coordinating external fertilisa-tion, 9, 91, 41–43
larval release, 77Agrotis ipsilon (black cutworm moth)
male response activated by juvenilehormone, 189
Alarm pheromonescommercial exploitation
for aphid control, 265hatchery trout taught to
recognise predator, 254costs to responding
aphid drop, 152costs to sender
eavesdropping by predators, 229, 234
evolution from pre-existing chemical cues, 9
evolution in related individuals,147–157
family groups, 147, 148, 149kin selection, 148–157
evolution in unrelated individuals,157–162
Diadema antillarum(sea urchin), 158
fish, 158–162predator labelling with host
pheromoneAeolidia papillosa (sea-slug), 148predatory fish, 161
propaganda by plants to repel aphids, 244, 245prey to escape ants, 244robber bees, 243slave-making ants, 241, 244
Alcelaphus buselaphus cokei (Coke’s hartebeest)
self marking and presentation, 92Aleochara curtula (rove beetle), 61Alkylmethoxypyrazine, 75Allee effects, 75
feeding, 79Dendraster excentricus
(sand dollar), 80mating, 79
external fertilisation, 79internal fertilisation, 79
overcoming tree defencesbark beetles, 81, 81
for wildlife conservation, 254Allelochemicals, 2Allomarking, 111Allomones, 2
deceit, propaganda, 230Alloparental care, 123
egg dumping, 230Alpheus heterochaelis (snapping shrimp)
modulation of visual threat signalsby pheromones, 18
Alternative mating strategiessatellite males
eavesdropping parasitoid pressure, 233
Nauphoeta cinerea (African cockroach), 60
she-malesAleochara curtula (rove beetle), 61Thamnophis sirtalis parietalis
(garter snake), 60Alzheimer’s disease, diagnosis, 297Amblyomma hebraeum (tick)
pest management, 262Amniotic fluid, 199Amphibians
Bufo borealis (western toad)tadpoles, alarm pheromone,
escape from predator, 157Litoria splendida (magnificent
tree frog)peptide sex pheromone, 21
tadpoles, kin, and alarm
371
Italic numbers refer to figures, boxes and tables
Organisms are nearly all indexed under taxonomic names in main entries; where commonnames are used a cross-reference is provided to the taxonomic name (except for groups).Taxonomic and/or common names are used in subentries.
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372 INDEX
Amphibians (cont.)pheromones, 157see also Salamanders and newts
Amphid sensilla, 203Amphiprion (anemone fish), 238, 239Amygdala
interaction between main olfactoryand vomeronasal organ (VNO)outputs, hamster, 185
Anal glandCastor canadensis (beaver)
variation between families, 96Meles meles (European badger), 111Mustela erminea (stoat)
repel prey, 265Anas platyrhynchos (mallard duck), 21Andrena nigroaenea (solitary bee)
duped by orchid, 240, 2423�-Androstenol, 253, 288, 289
steroid sex pheromones in pig, 20in truffles, use of sows to find, 253
5�-Androstenone, 191, 288, 289Boar Mate™, 252, 253, 254
Anguilla rostrata (American eel)zigzag behaviour, 227
Animal welfareunderstanding olfactory world, 254
Anomala osakana (Osaka beetle), 17Anonymous signals
social insects, 102Anosmias
induced changes on exposure, 191specific, 291
Antagonists, 66, 68, 187, 227, 260Antelopes
Kobus kob (Uganda kob)lekking, 58
Kobus leche (Kafue lechwe)lekking, 58
Litocranius walleri (gerenuk), 89Oreotragus oreotragus
(klipspringer antelope)scent marks eavedropped by
ticks, 233Ourebia ourebi (oribi), 87
border maintenance by multimale groups, 96
territories, 96Raphicerus melanotis (grysbok), 13
Antennal lobe, 165macroglomerular complex
(MGC), 165main, 165
Anthonomus grandis (boll weevil), 20pest management, 263
Anthopleura elegantissima (sea anemone)alarm pheromone (anthopleurine),
148, 149Anthopleurine, 148
soluble polar pheromone, 15Anti-aggregation pheromones, 235Antilocapra americana (pronghorn
antelope)alert signals, 148
Antorbital glandsterritorial and self marking
Alcelaphus buselaphus cokei(Coke’s hartebeest), 92
territorial marking by antelope, 89, 233
Aphaenogaster, see NovomessorAphrodisiac pheromones, see Male
pheromonesAphrodisin, 182Apis cerana japonica (Japanese
honeybee), 154–155, 156–157Apis mellifera, see HoneybeesAplysia (sea-slug)
peptide sex pheromone, 15Apocephalus paraponerae (phorid fly)
eavesdrops host pheromone, 233Apolipoprotein D (apoD), 290Aposematism
insects, aggregation, 75toad tadpoles, 157
Applications of pheromonesanimal husbandry
primer pheromones, 252signalling pheromones, 253
aquaculturehatchery trout taught to
recognise predator odour, 254induce sperm production in
carp, 254beneficial insects
honeybees, 255captive breeding rare species, 254
kin recognition, mate choice, 254commercialisation, 267–268
economics, 268public policy, 268
pest managementalarm pheromones, aphids, 265deterrent odours, 264–265lure and kill or mass trap,
260, 261–264
marine fouling organisms, 266mating disruption, 257–260,
261, 267monitoring, 256, 256, 257pest resistance to pheromones, 267primer pheromones, 265–266push-pull or stimulo-deterrent
diversionary strategies, 265, 266
self-protecting plants, 266pets
cats, spray marking, 254minipigs, 254
Tuber melanosporum (truffles), finding, 253
Aquatic pheromonesalarm pheromones
Diadema antillarum (sea urchin), 158
fish, 158toad tadpoles, 157
coordination of external fertilisationCarassius auratus (goldfish), 9, 19Nereis succinea (polychaete worm), 42
Dendraster excentricus (sand dollar)larval settlement, 80
Diadema antillarum (sea urchin)alarm responses, 158
eicosanoids (PUFAs)barnacle hatching pheromone, 77
Homarus americanus (lobster)precopulatory mate guarding,
61, 62peptides
Aplysia (sea-slug), 15Nereis succinea (polychaete
worm), 42sodefrin (newt), 15, 16
polar pheromoneanthopleurine, sea anemone,
15, 148, 149prostaglandins
Carassius auratus (goldfish), 19,27–28
Temora longicornis (copepod), 213see also Pheromones, aquatic habitats;
Signals, design, aquatic habitatsArachnids
mitesVarroa, 116, 262
phalangids (harvestmen)use ant alarm pheromone,
defence, 244
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INDEX 373
spidersbolas spiders, 240, 243Habronestes bradleyi, attracted to
fighting ants, 229Linyphia litogiosa, 59
ticksguanine, assembly pheromone, 80Ixodes neitzi, 233
Argyortaenia velutinana (moth)male pheromone response wider
than range females produce, 69Argyropelecus hemigymnus
(deep-sea hatchet fish), 207Arms race
bark beetles and their predators, 234
Arthropodin, see Settlement-inducingprotein complex (SIPC)
Artificial insemination (AI), 253Assembly pheromone
ticks, 80Assortative mating, 65Asymmetric tracking, 68Atemeles pubicollis (beetle social para-
site of Myrmica ants), 246Atta (leaf-cutter ants), 114, 263
Atta texanasensitivity of workers to trail
pheromone, 136
‘Badges’ of statusNauphoeta cinerea (African
cockroach), 44Bark beetles
push-pull or stimulo-deterrent diversionary strategies, 266
see also Aggregation pheromones;Allee effects; Arms race; Dendroctonus; Exploitation ofpheromones by organisms; Ips;Mice; Pheromones
Barnaclesdisrupting settlement, 266egg hatching pheromone, 77larval settlement, 26, 78
Beetle aggregation pheromones, 81Behavioural manipulation by semen
componentsDrosophila melanogaster, 63humans? 64Thamnophis sirtalis parietalis
(garter snake), 63Benzaldehyde, 5, 243
Bioassays, see Pheromone identifica-tion, bioassays
BirdsAnas platyrhynchos (mallard duck)
possible sex pheromone, 21eavesdropping vole scent marks by
ultraviolet cuesButeo lagopus (rough-legged
buzzard), 232Falco tinnunculus (kestrel), 232
glandsanal, 21sebaceous, 21uropygidial, 21
nest recognition, 21pheromones, 21procellariiform seabirds
olfaction, 21zigzag flight upwind, 225
cis-(Z)-�-Bisbolene epoxide, 233Boar Mate™, 252, 253, 254Boiga irregularis (brown tree snake)
pest management, 262Bombykol
identification of, 25Bombus, see BumblebeesBombyx mori (silk moth), 23
antennae, design, 166Bos taurus (bulls and cows), 253Boundary layer
effects on olfactory sensors, 174turbulence, 219
Brain, see Olfactionendo-Brevicomin, 235exo-Brevicomin, 5Bruce effect
memory and pregnancy block inmice, 201
pregnancy block by malepheromones, 195
Bufo borealis (western toad)tadpoles, alarm pheromone, 157
Bumblebees (Bombus)marking visited flowers, 85
Butenandt, Adolf, 23Buteo lagopus (rough-legged
buzzard), 2322-sec-Butyl-dihydrothiazole
(thiazole or SBT), 193, 194
C20 polyunsaturated fatty acids (PUFAs) (eicosanoids), 77
Caenorhabditis elegans (nematode), 203
Callinectes sapidus, (blue crab)rheotaxis, 224
Callithrix jacchus (common marmoset)subordinate female ovulation
suppressed, 197Camponotus socius (ant)
group recruitment, 134Canis
Canis latrans (coyote)urine repells rodents, 265
Canis lupus (grey wolf)border marks in territory, 97suppression of subordinate
reproduction, 124Canis lupus familiaris, see DogsCanis sinensis (Ethiopian wolf)
border marks in territory, 97Carassius auratus (goldfish)
primer and releaser pheromones, 19
sex pheromones evolved from hormones, 9, 27–28
Castesdifferences
pheromone secretions, ants, 114responses to pheromones, social
aphids, 153social parasites copy odour badges
of termite host castes, 248see also Honeybees
Castor canadensis (beaver)anal glands, 96avoid marks at low population
densities, 91castoreum, 96kin recognition, 111pest management, 265suppression of subordinate
reproduction, 124Castoreum, 96Central-place foragers
naked mole rat 133social insects, 133
Ceratitis capitata (Mediterranean fruit fly)
leks, 58, 59eavesdropped by predator, 232
Ceratovacuna lanigera (social aphid)soldiers respond, non-soldiers
flee, 153soldiers attack predator, 153
Cerocebus albigena (mangabey,primate), 271
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374 INDEX
c-fos, 182Chemical camouflage, 247–248Chemical mimicry, 248–249Chemical structure, 302Chemoreception, 164Chirality, 3055�-Cholestane-3,24-dione
trail pheromone, Malacosoma, 130Chrysopa slossonae (lacewing larvae)
escapes detection by chemical camouflage, 247
cis (in chemical name), 305Citral, 243Clean-up enzymes, 177Clethrionomys glareolus (bank vole)
females prefer odours of dominantmale, 46
Cloacal gland, 16Coccinella septempunctata (ladybird
beetle)aposomatic pheromones, 75
Colony defencecoordinated response to
danger, 155Colony-level selection
honeybees, 107Colophina monstrifica (social
aphid), 153Combinatorial coding
glomeruli, 170Communal nesting
enabled by synchronous breeding, 196
mouse preference for kin, 54, 55Communication, 2, 3
privacy in, 16sex communication channel, 66see also Composite signals;
Pheromones; SignalsComposite (combined) signals
chirp and odours, naked mole ratforagers, 136
parallel sensory channels, 17pheromone and stridulation
recruitmentAphaenogaster (Novomessor)
(ant), 140Conflict between the sexes
Linyphia litogiosa (spider), 59Nicrophorus (burying beetle), 59semen components
Drosophila melanogaster, 63humans, 64
Thamnophis sirtalis parietalis(garter snake), 63–64
Conscious and unconscious responsesto odours, 185
human responses, 185electroencephalogram (EEG), 185functional magnetic resonance
imaging (fMRI), 185Contact chemoreception
ants detecting colony identity, 4Contests, 44–46Coolidge effect, 57, 202
Gryllus bimaculatus (cricket), 57male rodents, 57Vipera berus (adder), 57
Cooperative breeders, 123–127plural breeders, 192
subordinate female ovulationsuppressed, 124, 126, 127,197–198
singular breeders, 124Cooperative signal, 122Coptotermes (termite), 130Coremata, 10, 11
honest signal, 3sexual selection, 47, 49
Cornicle secretionsaphids, 150
Corpora allata, 190termites, 119
Corynebacteriumbacteria in human armpits, 291
Costs (of signalling)eavesdropping by parasites
Ixodes neitzi (ticks), 233parasitoid insects, 232
eavesdropping by predators, 231ant-decapitating flies, 234lekking Ceratitis capitata (Mediter-
ranean fruit fly), 232spiders attracted to fighting
ants, 229low metabolic cost of pheromones
Anthonomus grandis (boll weevil), 20
Litoria splendida (magnificent treefrog), 21
long-distance communication,207
riskinvestigating marks, beavers, 89
timemarking, antelope, 87
Countermarking, see Over-markingCourtship
Cerocebus albigena (mangabey,primate), 271
Creatonotus gangis (arctiid moth), 10Drosophila melanogaster (fruit fly),
18, 70Grapholitha molesta (oriental fruit
moth), 7Homarus americanus (lobster), 61, 62humans, 277Macaca mulatta (rhesus monkey), 281newts, 16salamanders , pheromone delivery
patterns, 4species recognition, 64, 70, 72
Crataegus mollis (hawthorn), 85Creatonotus gangis (arctiid moth)
coremata, 3, 10Cremogaster dohrni artifex (attendant
ant of butterfly), 238Crocuta crocuta (hyena)
expose scent glands on approach, 91Cross-fostering experiments
Acomys cahirinus (spiny mouse)diet cues, 110
Odocoileus hemionus hemionus (black-tailed deer), 199
honeybees, 106MHC preferences of mice, 55Spermophilus beldingi (Belding’s
ground squirrel), 110Crotalaria
plant source of pyrrolizidine alka-loids (PAs), 47
CrustaceansAlpheus heterochaelis (snapping
shrimp)modulation of visual threat by
pheromones, 18barnacles
larval settlement, 26Semibalanus balanoides (barnacle),
77settlement behaviour, 78
copepodsLepeophtheirus salmonis (salmon ec-
toparasite), control, 262crabs
Callinectes sapidus (blue crab), 224Rhitropanopeus harrisii (mud crab),
77Homarus americanus (lobster)
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contest, 44precopulatory mate guarding,
61, 62tropotaxis, 222
Panulirus interruptus (Californiaspiny lobster)
aggregation for defence, 75plankton
Temora longicornis (copepod), 213Culex (mosquito), 263
oviposition pheromone, 75Cuticular hydrocarbons
ant trail pheromones, 140colony recognion
honeybees, 106ants, 108deception, 247–249Polistes fuscatus (paper wasp), 107
deception, 61dipteran sex pheromones, 70dipteran speciation, 71Gryllus bimaculatus (cricket)
mate choice, 51, 57solid phase microextraction (SPME)
from living ants, 33Cydia pomonella (codling moth)
pest management, 258Cynictis penicillata (yellow mongoose)
scent marking, 90Cynops pyrrhogaster (red-bellied newt)
courtship, 16sodefrin, 15, 16
Cypris larva (barnacle), 78L-Cysteine-glutathione disulfide, 42
Danaidone, 4Danaus gilippus (queen butterfly)
transfer pheromone direct to female, 4
Darwin, Charlessexual selection on pheromones, 37
Dauer larvaCaenorhabditis elegans
(nematode), 203Dear-enemy phenomenon (territorial
defence), 99, 112Deer
Odocoileus hemionus columbianus(black-tailed deer)
alert signals, 28, 147–148composite (combined) alarm
signal, 17scent marking, 91
2,3-Dehydro-exo-brevicomin (brevicomin or DHB), 5, 193, 194
2,5-Demethylpyrazine, 194Dendraster excentricus (sand dollar), 80Dendroctonus (bark beetles), 81
Dendroctonus brevicomiseavesdropped by predator, 234
Dendroctonus frontalis, 82male inhibition by male
pheromones, 235Dendroctonus micans, 81, 81Dendroctonus montanus, 81Dendroctonus pseudotsugae, 82Dendroctonus valens, 81
Dermacentor variabilis (tick)pest management, 262
Desmognathus ochrophaeus (salamander)male injects pheromone into
female, 4Development
agehamster, 189honeybee worker responses, 190honeybee, queen mandibular
pheromone (QMP), 121, 122Agrotis ipsilon (black cutworm
moth), 189caste differences in social insects
pheromone production andperception, 190
learningmale hormone surge to vaginal
odours (hamster), 182maternal behaviour, 191, 200Rhagolitis pomonella (apple maggot
fly), 190–191social insects, 107, 248
Schistocerca gregaria (desert locust), 76
sensitive periodewe learning odour of her lamb,
199, 200kin recognition, 110newly adult ant, 248
sex differences in brainmammals, 190moths, 190
switchdauer pheromone, 203social insects, 119, 204, 204
Diadema antillarum (sea urchin)alarm responses, 158
Diastereoisomers, 307Diet
effect on odourAcomys cahirinus (spiny
mouse), 110humans, 274
quality and mate choiceMicrotus pennsylvanicus
(meadow vole), 46Utetheisa ornatrix (tiger moth)
hydroxydanaidal (HD), 472,5-Dimethylpyrazine
female mice, Lee-Boot effect, 195N,N-Dimethyluracil, 130Dinoponera quadriceps (ant)
cuticular hydrocarbons, non-destructive study by solidphase microextraction (SPME),33–34
Diomedea (albatross)navigation, food location, 21, 225
Dipsosaurus dorsalis (iguana)composite (combined) signal with
ultraviolet cue, 14Disassortative mating
MHC genotype, 55(Z,Z,E)-Dodecatrienol, 130, 140(Z)-7-Dodecen-1-yl acetate, 1Dogs (Canis lupus familiaris)
ability to track odour trail, anddetect its direction, 218
as detector of cow oestrus, 253distinguishing humans by
smell, 274forensic use of smell, 296olfactory sensory sensitivity, 167scent marking landmarks, 99
Dolichotis patagonum (mara)males directly mark females, 100
DominanceHomarus americanus (lobster), 44, 61mice, 39, 93, 193, 194Microcebus murinus (lesser mouse
lemur), 196Microtus pennsylvanicus (meadow
vole), 46Nauphoeta cinerea (African
cockroach), 40, 44, 45queen, social insects, 118in queenless ants, 34–34social mammals, 124
Drakaea glyptodon (orchid) dupeswasp, 241
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Drosophila, 36, 71, 188, 209Drosophila adiastola subgroup, 72Drosophila mauritiana, 72Drosophila melanogaster, 63, 72
conflict between the sexes, 63courtship, pheromones, and
speciation, 18, 70, 71genetic engineering, 35–36, 70
Drosophila sechellia, 72Drosophila simulans, 71
Dufour’s gland, 11, 130ant trail, 139Lasioglossum zephyrum (sweatbee), 108parasitoid wasps, 84, 84queen bee egg mark, 122slave-making ants, 241, 244
Eavesdropping, see Exploitation ofpheromones by organisms
EchinodermsDendraster excentricus (sand dollar)
larval settlement, feedingbenefit, 80
Diadema antillarum (sea urchin)alarm responses, 158
Eciton (army ants), foraging, 141–143Egg dumping, 230
fish, 231insects, 230
Egg hatching pheromoneSemibalanus balanoides (barnacle), 77
Eicosanoids (PUFAs) (prostaglandinsin vertebrates)
barnacle hatching pheromone, 77Electroantennogram (EAG), 29, 30Electroencephalogram (EEG)
response to odours, 185Electroolfactogram (EOG), 189Electrovomerogram (EVG), 194Elephants
African (Loxodonta africana), 47mate guarding, 62
Asian (Elephas maximus), 1, 5, 47musth, testosterone elevation, 48pheromone shared with moths, 2
musthfemale response, 47male aggressiveness, 47temporal gland secretion, 47, 48testosterone elevation, 47, 48urine, 47
young males smell different fromadults, 47
Elephas maximus, see Elephants, AsianEnantiomers, 17, 305Enemy specification, 155Enoclerus lecontei
eavesdropping, 234, 236Entgegen (E), opposite, 308Environmental odour cues and
recognition, 103, 110Epiphyas postvittana (light brown apple
moth)pest management, 257, 258, 259
EphestiaEphestia elutella (tobacco moth), 47Ephestia kuehniella (flour moth)
pest management, 263Equus (horse), flehmen to oestrous
urine, 179Esox lucius (pike), 254
labelled with prey alarmpheromone, 159–161
naïve prey learn odour, 159predator attraction, interference
hypothesis, 162, 162Estrus, see Hormones; OestrusEstrogen, see Hormones; Oestrogen4-Ethyl-2-methoxyphenol, 44Ethyl-trans-cinnamate, 7Euglossine bees
courtship needs plant perfumeoils, 12
Eupoecilia ambiguella (grape moth)pest management, 258
Eusociality, 116conflict over reproduction in
societies, 118continuum (reproductive skew),
117, 117social insects and social mammals,
parallels, 127see also Social insects; Social
mammalsEvolution, see Pheromones,
evolutionExperience, see Development;
LearningExperimental methods, see
Pheromone identificationExploitation of pheromones by
organismsarms race, 230code breaking, 229deception
bolas spiders, 240, 243
cost of excluding deceivers, 249pollination by sexual deception,
240, 241, 242predators, guests and parasites of
social insects, 244–249propaganda, 241–244, 245
deception by mimic chemicalsenabled by shared biochemistry
in all life, 230eavesdropping, 229
aggregation pheromones, 81, 234,235, 236
alarm pheromones, 229, 234egg dumping, 230–232enabled by characteristics of
olfaction, 230intraspecific, sex pheromones as
aggregation pheromones, 236prey responses to predator
pheromones, 154–155, 156–157,236, 243, 264–265
sex pheromones, 231–232territorial marking
pheromones, 233intraspecific
hormones evolving intopheromones, 8
mutualism, 237‘communication’ between leaf-
cutter ants and fungus, 238ants and lycaenid butterfly
caterpillars, 237aphids tended by ants, 237sea anemones and anemone fish,
238–240External fertilisation
coordination by pheromones, 41Carassius auratus (goldfish), 9, 19Nereis succinea (polychaete
worm), 42
Fabré, Jean-Henri, 23Faeces, territorial marking
antelopes (oribi), 96Meles meles (European badger), 98salamanders, 95
Falco tinnunculus (kestrel), 232Farnesene, 130, 193, 194
produced by wild potato, 244, 245Feliway™ (synthetic cheek gland
pheromone), 254Female choice, see Mate choice; Sexual
selection
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Female pheromoneseffects on females
biases sex ratio in pregnancy, 197delayed puberty, 194, 195oestrus synchronisation, 196, 282social insects, 115–116, 119, 121, 123subordinate female ovulation
suppressed, 197suppressed oestrus cycles, 194, 195
effects on malesMicrocebus murinus (lesser mouse
lemur), 196Femoral gland
Iguana iguana (lizard)testosterone influence, 46
Fishalarm pheromones, 158–162Amphiprion (anemone fish
symbiotic with sea anemones),238, 239
Anguilla rostrata (American eel)zigzag behaviour, 227
Argyropelecus hemigymnus (deep-seahatchet fish)
biggest nasal organ for body size,vertebrate, 207
Carassius auratus (goldfish)pheromones from hormones, 9,
19, 27–28, 43, 189Esox lucius (pike)
labelled with prey alarmpheromone, 159–161
naïve prey learn odour, 159odour to teach hatchery rainbow
trout, 254predator attraction, interference
hypothesis, 162, 162Gadus morhua (cod)
zigzag orientation in plume, 225Gasterosteus aculeatus (stickleback)
MHC and mate choice, 55Lepomis cyanellus (green sunfish)
alloparental carer, 230–231Notropis umbratili (redfin shiner)
interspecific egg dumper, 231Oncorhynchus mykiss (rainbow trout)
taught to recognise predatorodour, 254
Petromyzon marinus (marine lamprey)control, 262
Phoxinus phoxinus (Europeanminnow)
alarm pheromone, 158
Plotsus lineatus (catfish)scanning electron microscopy of
olfactory epithelium, 169Premnas (anemone fish symbiotic
with sea anemones), 238Primiphales promelas (fathead min-
now), 158Puntius (androgen effects on elec-
troolfactogram (EOG)), 189salmon
zigzag orientation in plume, 225Flame ionisation detector (FID), 30Flehmen
Bos taurus (bulls and cows), 253Elephas maximus (Asian elephant), 48Equus (horse), to oestrous urine, 179
Flicks, antennalbehaviour of smelling, 174Homarus americanus (lobster), 174
Floaters, 99Fluctuating asymmetry, 49
humans, 280Panorpa japonica (scorpion fly), 49
Formica subintegra (slave-making ant),241, 244
Formicoxenus (shampoo ant)social parasite of ants (Myrmica),
247Frequency dependant responses
parasitoid wasp superparasitism,188
Fright response, 158–159Functional groups, 302Functional magnetic resonance
imaging (fMRI)humans, 174, 175, 185
Gadus morhua (cod)zigzag orientation in plume, 225
Galago (bushbaby)nocturnal, use of pheromones, 13
Galaxolide, 185Gargaphia solani (aubergine lace bug)
intraspecific egg dumping, 230subsocial care, alarm pheromone,
147Garter snake, see Thamnophis sirtalis
parietalisGas liquid chromatography (GC), see
Pheromone identification, separating chemicals
Gasterosteus aculeatus (stickleback), 55Genetics
Caenorhabditis elegans (nematode)using mutants, 203
Drosophila, 35–36, 70humans, perception of odours,
specific anosmias, 291–292moths, pheromone signal, 69olfactory receptor proteins (OR), 169
Geometrical isomersEntgegen (E), Zusammen (Z), 308
Geraniolalarm pheromone
Gargaphia solani (aubergine lacebug), 147
Gestalt modelcolony odour, 108
Glands, 9, 10, 11adrenal
female mice, Lee–Boot effect, 195anal
birds, 21Castor canadensis (beaver), 96Meles meles (European badger),
111Mustela erminea (stoat), 265
antorbitalterritorial marking by antelope
(gerenuk) 89, 233cloacal, 16closable, due to eavesdropping
predators, 233Dufour’s
ant trail, 139Lasioglossum zephyrum (sweatbee),
108parasitoid wasps, 84, 84queen bee egg mark, 122slave-making ants, 241
femoralIguana iguana (lizard), 46
labialtermites, 138
mandibularTrigona (stingless bee), 136
musk, 148Nasonov
honeybees, 131poison
ant, 131postcloacal
salamander, 95postpharyngeal
ants, 108, 248preorbital, see antorbital
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378 INDEX
Glands, (cont.)preputial
mice, 193pygidial
ant, 134sebaceous
birds, 21humans, 284signal life, 14
some vary more within aspecies, 113
sternalant, 130Malacosoma americanum (tent
caterpillar), 130Polistes (wasp), 107termite, 130, 137
subcaudalMeles meles (European badger), 111
sweatapocrine, 284eccrine, 284
temporalelephants, 46
uropygidialbirds, 21
Glomerulicombinatorial coding, 170–174functional units of processing, 172
L-Glutamic acid, 42L-Glutamine, 42Gossyplure, 303G-proteins, 166Grandlure, 263Grapholitha molesta (oriental fruit moth)
courtship display of hair pencils, 7pest management, 258response to pheromone blend, 67
‘Green beard’ phenomenon, 104, 105Gryllus bimaculatus (cricket)
Coolidge effect, 57mate choice for optimal
outbreeding, 51Guanine, 80
Habronestes bradleyi (spider), attractedto fighting ants, 229
Hair pencilsmoth, 7
Hamilton’s ‘selfish-herd’ theoryfish shoals and alarm
pheromone, 159Hamilton’s rule, kin selection, 116
Hawaiian Drosophila fruit flies, 72Drosophila adiastola subgroup, 72speciation, 72
Helicoverpa zea (corn ear wormmoth), 187
Helogale undulata (dwarf mongoose)individual recognition, 112
(Z)-7-Heneicosene, 61Heneicosane, 842-Heptanone, 194Herpestes auropunctatus (Indian
mongoose)individual recognition, 112
Heterocephalus glaber (naked molerat), 125
colony odour, parallels with socialinsects, 112
queen suppression of worker reproduction, 124
recruitment of foragers, 136Heterodera glycines (soybean cyst
nematode)pest management, 258, 260
E-10,Z-12-Hexadecandien-1-ol, 25Z-9-Hexadecenal, 130Hexanoic acid, 130Hinterland marking, 97–98Homarus americanus (lobster), 44
antennal flicks, 174precopulatory mate guarding, 61, 62tropotaxis, 222
Homo sapiens, see HumansHonest signal, 2, 44, 45
queen pheromonehoneybees, 121, 121
reflecting biological state ofmarker
territorial scent marks, 89Utetheisa ornatrix (tiger moth), 47,
49, 49, 50Honeybees (Apis mellifera)
Africanised bees, monitoringspread of, 256, 256
cell capping, odour cues, 116colony odour sources, 106colony-level signature, 106comb wax odour cues, 106manipulating behaviour for bee-
keepingdelaying swarming, 255directing pollination, 255luring swarms, 255
marking visited flowers, 85
multiple paternity in nestblending of template colony
odour, 105Nasonov pheromone
marking new nesting cavity, 131swarming, 131
pheromones, summary, 115queen mandibular pheromone
(QMP), 121as honest signal, 121as sex pheromone, 19rapid colony response to loss, 121transmission, 121
queen-rearing nepotism, 107queen retinue, 116sting pheromone attracts other
bees to sting, 154swarming, 121Varroa mite, 116, 262
Hormonescortisol and recognition of human
babies, 276humans
birth, 201cortisol and recognition of
human babies, 276juvenile hormone ( JH)
Agrotis ipsilon (black cutwormmoth), 189–190
social insects, 204, 204oestrogen, 198oxytocin in ewes, 199, 200, 201pheromone influence on hormones
female stimulates testosterone secretion in lesser mouselemur, 196
male hormone surge to vaginalodours in hamster, 182
puberty acceleration in rodents,ungulates and non-human primates, 193
responses induced via olfactorysystem, 185
sexual activity in Petaurus breviceps (sugar glider), 196
testosteroneandrogen effects on electroolfac-
togram, fish, 189femoral gland, Iguana iguana
(lizard), 46graded response of scent glands
in Microtus pennsylvanicus(meadow vole), 46
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hamster male brain needs to respond, 189
male sexual response in mammals, 183
musth and temporal gland secretion in elephants, 47, 48
see also Primer pheromones; Female pheromones; Malepheromones
Host-discrimination, 83Host-marking pheromones (HMP), 74,
83–85evolution of, 83host-discrimination, 83individual recognition, 85
Human leucocyte antigen (HLA), seeHumans, mate choice; Majorhistocompatibility complex;Mammals, mate choice
Humansadvertisement of oestrus, 281
‘copulins’, 281unresolved in humans, 282
age, changes withperception, 271secretions, 275sensitivity, 297
anosmiasinduced changes on exposure,
191specific, 291
antiperspirants and deodorants,288
axilla (armpit) hair and odour creating bacteria, 288
behavioural manipulation by semen hormones? 64
candidate compounds, 285axillae (armpits), 287–291glands, 284sites over body, 286, 287
cleanliness not necessarily avirtue, 276
cultural and social aspects ofodours, 273
osmologies, 273Napoleon, 273sexual attraction, 273
importance of odours in humanbehaviour and biology, 274
mate choice, 2775�-androstenone, 277body symmetry and, 280
kibbutz or Westermarck effect,280
human leucocyte antigen (HLA),52, 278
major histocompatibilitycomplex (MHC), 52–56,278–280
sexually selected traits, 277mothers and newborn babies, 275
correlation of mother cortisoland baby recognition, 276
importance of natural odours,275
speed of learning, 275odour preferences
learning, 271neonates different from older
children and adults, 271odours and memory
experimental tests, 276Proust, 276
olfactionmapping brain activity, 174, 175,
185olfactory receptors
genes and pseudogenes, 170perception of odours, 291
changes during menstrual cycle,292, 293
changes with puberty, 292specific anosmias, 291, 292variation between individuals,
291perfumes, 273, 279phenotype matching, 276primer pheromones
age of female puberty, 285menstrual synchrony, 282–285steroids and mood, 295
recognition, 274children, sibs, parents, 275security blankets, 276self, partners, 275T-shirt sniff tests, 274
semen components, 64truffles, 253, 288using human odours, 295
‘human pheromones’, 298forensics, 296medical diagnosis, 296–297mood changers, 298
vomeronasal organ (VNO),putative, 294
genetic evidence, 295location, 294neuroanatomical and histologi-
cal evidence, 295see also Primates
HutteritesMHC and partner choice, 280
Hydrobates pelagicus (storm petrel)nest recognition, 21
Hydrocarbons, see Cuticularhydrocarbons
Hydroides (polychaete worm)plankton settlement, 79
3-Hydroxy-2-butanone, 446-Hydroxy-6-methyl-3-heptanone,
193, 194Hydroxydanaidal (HD), 50
derived from plant pyrrololizidinealkaloids in diet, 47
Hypothalamus, 184Microtus ochrogaster (prairie vole), 198oxytocin
maternal behaviour, 199, 200Hypoxanthine 3-N-oxide, 158
Iguana iguana (lizard)forest habitat, volatile pheromone,
14–15femoral gland
testosterone influence, 46Imprinting
mate recognition, 198odours, recognising kin and colony
members, 199Individual recognition
generalisation across glandodours, 113
Helogale undulata (dwarf mongoose),112
Herpestes auropunctatus (Indianmongoose), 112
Homarus americanus (lobster), 46humans, 274–275mechanisms, within a species some
glands vary more than others,113
Meles meles (European badgers), 112mice, 112pair bonding
Niveoscincus microlepidotus(lizard), 113
Proteles cristatus (aardwolf)‘dear-enemy’ recognition, 113
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380 INDEX
Information centreMalacosoma americanum (tent
caterpillar), 133Inosine, 42Integrated pest management
(IPM), 255pheromones in control of tomato
pests, 259Ips (bark beetles)
Ips paraconfusus, 235discovery of synergy, 35
Ips pinibioassays, 24
Ips typographuspest management, 264
(R)-(�)-ipsdienol, 234(S)-(�)-ipsdienol, 234Iridomyrmex (ants), 130Isomers, 304
constitutionalfunctional group, 304positional, 304
naming of, 305stereoisomers
chirality and enantiomers (optical isomers), 305–307
Ixodes neitzi (tick)responds to scent marks of host
antelope, 233
(R)- and (S)-Japonilure enantiomersuse by sympatric beetles, 17
Juvenile hormone (JH)Agrotis ipsilon (black cutworm moth)
male response activated by, 189social insects, 204
ants, 204termites, 119
Kairomones, 2eavesdropping, 230
Kangaroosmate guarding by male, 62
Keifera lycopersicella (tomato pin worm)pest management, 258, 259, 261
Kibbutz or Westermarck effecthuman mate choice, 280
Kin recognition 103–105mate choice
conservation projects, 254mechanisms, 103
low metabolic costs, 103Kin selection, Hamilton’s rule, 116
KobusKobus kob (Uganda kob), 58Kobus leche(Kafue lechwe), 58
Labelled lines, 187Labial gland
termites, 138Lasioderma serricorne (cigarette beetle)
aggregation pheromone, 83, 84Lasioglossum zephyrum (sweatbee)
guard bees discriminate relatedness, 108
Lasius (ants)trail pheromone, 130Lasius alienus
alarm pheromone, causes panicand escape, 155
Lasius fuliginosustrail following, 216
Lasius nigerforaging effort, food value, 139
Lateral olfactory tract, 183Learning
fish, conditioned alarm response topredator odours, 159, 191, 254
mammalsinduced peripheral changes in
response to odours after contact, 191
maternal behaviour, 191needed to distinguish oestrous
from dioestrous female odours,191
maternalewes bond quicker on subsequent
births, 200odour stimulation and
noradrenaline release inolfactory bulbs, 199
Rhagolitis pomonella (apple maggotfly), 190–191
see also DevelopmentLee–Boot effect, 194, 195Leks, 57
Ceratitis capitata (Mediterraneanfruit fly), 58
cost, eavesdropped by predators,231–232
Kobus kob (Uganda kob), 58Kobus leche (Kafue lechwe), 58Lutzomyia longipalpis (sandfly), 58Xylocopa fimbriata (carpenter bee),
57
Lemur catta (ring-tailed lemur)group territorial ‘stink fights’, 90
Leontopithecus rosalia (lion tamarin)suppression of subordinate
reproduction, 127Lepeophtheirus salmonis (salmon
ectoparasite)pest management, 262
Lepomis cyanellus (green sunfish)alloparental carer, 230–231
Leptothorax acervorum (ant)tandem running recruitment, 134
Leptotyphlops dulcis (blind snake), 247Lestrimelitta limao (robber bee), 243Linoleic acid, 84Linyphia litogiosa (spider)
conflict between the sexes, 59Liriomyza sativae (leafminer fly), 259Litocranius walleri (gerenuk), 89Litoria splendida (magnificent tree
frog)peptide sex pheromone, 21
Lobesia botrana (grape vine moth)pest management, 258
Lordosis, 20, 184, 253Loxodonta africana, see Elephants,
AfricanLuteinising hormone (LH), 183
Microtus ochrogaster (prairie vole),198
Luteinising hormone-releasing hormone (LHRH), 198
Lutzomyia longipalpis (sandfly), 263lekking, 58
Macaca mulatta (rhesus monkey), 281Macroglomerular complex (MGC), 165Macrotermes (termite)
nest building, 143–144Maculinea rebeli (lycaenid butterfly)
manipulates ant host, 249Main olfactory bulb (MOB), 165Main olfactory epithelium (MOE), 165,
178, 179, 182, 183Main olfactory system
detection of pheromones, 181Oryctolagus cuniculus (European
rabbit), 181Sus scrofa (pig), 184
sheep maternal behaviour, 201see also Olfaction
Major histocompatibility complex(MHC)
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bacterial flora and odours, 53Bruce effect, 195communal nesting, mouse
preference for kin, 54, 55familial imprinting
odour preferences for differencein mates, 55
human leucocyte antigen (HLA),52, 278
kin recognition in other vertebrates(fish, birds, reptiles), 56
mate choice, 52MHC diversity, Gasterosteus
aculeatus (stickleback), 55mice and humans, 54mutual distinguishing MHC of
odours inrats, mice and humans, 52
not universal, 56odours as marker for degree of
kinship, 52semi-natural experiments, 52
preferenceslearning, 55women taking oral
contraceptives, 54selection pressures maintaining
diversity of MHC alleles, 55, 56disassortative mating, 55higher implantation rates if dis-
similar, 55reduced chance of inbreeding if
dissimilar, 55source of the odours, 52–54
Major urinary proteins (MUPs),194, 195
longevity of signal, slow release,193
elephants, 195mice, 195
Malacosoma (tent caterpillars)trail pheromone, 130Malacosoma americanum, 133
laying trail, 135Male pheromones
diverse release structures, 7, 10, 70effects on females
leks, 57–59mice, Bruce effect (pregnancy
block), 195oestrus induction and puberty
acceleration (Whitten and Vandenbergh effects), 192
oestrus synchronisation, mice,194
effects on males, 196sexual activity of subordinates
suppressed, 196selection by female to avoid hybrid
matings, 70Mammals
group, kin, family, and individualrecognition, 109
individual recognition, 112Helogale undulata (dwarf
mongoose), 112Herpestes auropunctatus (Indian
mongoose), 112Meles meles (European badger), 112mice, 112
kin recognitionAcomys cahirinus (spiny mouse),
110mate choice
major histocompatibility complex (MHC), 52–56
mother–infant recognition, 109scent marking, 87scent sources, summary, 10secretory glands, 11see also Humans; Primates; Social
mammalsMan, see HumansMandibular gland
Trigona (stingless bee), 136Manduca sexta (tobacco horn moth)
zigzag orientation upwind, 225Marine invertebrates, alarm
responses, 158Marking enemies for further attack,
155Marsupials
kangaroosmate guarding by male, 62
Monodelphis (opossum)vomeronasal organ (VNO) needed
for oestrus induction, 181Petaurus breviceps (sugar glider)
scent marks affect sexual activity, 196
MastophoraMastophora bisaccata (bolas spider),
243Mastophora hutchinsoni (bolas
spider), 240Mate choice
avoiding lethal alleles, ‘t-complex’in mice, 56
diet qualityMicrotus pennsylvanicus (meadow
vole), 46fluctuating asymmetry
humans, 280Panorpa japonica (scorpion fly), 49
MHC diversity, see Major histocompatibility complex,mate choice
optimal outbreeding, 51Gryllus bimaculatus (cricket), 51Lasioglossum zephyrum (sweat-
bees), 51parasites, 51
mice, preference for uninfectedmates, 51
paternal investment and femalechoice, 47
Utetheisa ornatrix (tiger moth), 47pheromones
Ephestia elutella (tobacco moth), 47Mate guarding, see Sperm competition
and mate guardingMaternal behaviour and recognition
humans, 275sheep, 199
Mating disruption, as pest control,258
Medial amygdaloid nucleus, 183Medial preoptic area (MPOA), 182, 183Megaponera (ant), 130Meles meles (European badger)
anal glands, 111border and hinterland latrines,
98, 98individual recognition, 112subcaudal glands, 111
Memory and pregnancy block (Bruceeffect) in mice, 201
accessory olfactory bulb (AOB), 201Mesocricetus auratus (hamster) , 183
aphrodisin, 182over-marking, 100self-matching, 105
Messor barbarus (harvester ant)matching trail patterns to food
distribution, 1394-Methyl-3-heptanone, 2346-Methyl-5-hepten-2-one, 243(E)-3-Methyl-2-hexenoic acid ((E)-3M2H
or TMHA), 289
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Methyl-4-methylpyrrole-2-carboxylate,130, 136
2-Methyliazolidine, 44MHC, see Major histocompatibility
complexMice (Mus musculus domesticus)
Bruce effectmemory and pregnancy block,
201individual recognition, 112Lee–Boot effect, 195male pheromones
oestrus induction and puberty acceleration in females (Whitten and Vandenbergh effects), 192
marking rates, dominants and subordinates, 94, 94
mate choicepreference for uninfected
mates, 51mate choice to avoid lethal alleles,
‘t-complex’, 56pheromone marks
importance for animal wel-fare, 254
scent marking by dominantmale, 93
signal sites, urine posts, 94similar pheromone to bark beetle, 5‘t-complex’, 56Vandenbergh effect, 192Whitten effect, 192
Microcebus murinus (lesser mouselemur)
female urine stimulates spermatogenesis and testosterone secretion, 196
pheromones and sex ratios, 197sexual activity of subordinates
suppressed, 196Microdon mutabilis (syrphid fly)
matches its ant host, 248Microtus
Microtus ochrogaster (prairie vole),198
unfamiliar male odours, stimulation of oestrus, 198
Microtus pennsylvanicus (meadowvole)
suppression of subordinate reproduction, 124, 126,197–198
graded response of scent glandsto testosterone implants, 46
Microtus pinetorum (pine vole)suppression of subordinate
reproduction, 127Miriamide, 84Mitral/tufted cells, 171, 172
Bruce effect (memory of mate), 201memory for lamb, 200
MolluscsAplysia (sea-slug), peptide sex
pheromone, 15snails, direction of a trail, 217
Monodelphis (opossum)vomeronasal organ (VNO) needed
for induction of oestrus, 181Monogamous partner recognition,
202Monozygotic twins
human, dogs have difficulty distinguishing, 274
sheep, lamb recognition experiment, 109
Mother–infant recognitionhumans, 275sheep, 109
Mouse, see MiceMulti-component pheromones, 187
alarm pheromones in socialinsects, 154
ant, alarm pheromone signal,212, 214
asymmetric tracking, 68major and minor components, 66male response wider than range
females produce, 69response is to whole blend, moths,
67specificity, 17, 66synergy
insects, 35mammals, 195
Musca domestica (house fly)pest management, 263
Mus musculus domesticus, see MiceMushroom body, 165Musk, 185Musk gland, 148Mustela
Mustela erminea (stoat), anal glands,265
Mustela putorius furo (ferret), 190Musth, see Elephants
Myrmecaphodius excavaticollis (beetle)parasite of Solenopsis (fire ant), 247
Myrmecophilous invertebrates, 245Myrmica (ants), 247
Myrmica sabuletimatching foraging effort to food
value, 139Myzus persicae (aphid)
repelled by alarm pheromone fromplant, 244
Naming pheromonesfunctional groups
prefixes and suffixes, 303Napoleon, 273Nasonov pheromone
honeybees, 131, 255Nasutitermes (termite), 120, 155
marking enemies for further attack, 155
recruitment pheromones, 140Nauphoeta cinerea (African cockroach)
‘badges’ of status, 44honest signal, 44, 45male contest and female choice,
40, 44satellite males, 60
NematodesCaenorhabditis elegans (model
system), 203Heterodera glycines (soybean cyst
nematode)mating disruption with sex
pheromone, 258, 260number of olfactory receptor
types, 172Nemeritis (parasitoid wasp)
host marking pheromone, 84(E,E,E)-Neocembrene-A
(E-6-cembrene A), 140Nereis succinea (polychaete worm)
pheromones coordinate externalfertilisation, 42
Newts, see Salamanders and newtsNezara viridula (southern green stink
bug)sex pheromone blend change due
to parasitoid, 233Nicrophorus (burying beetle), 59Niveoscincus microlepidotus (lizard)
individual recognitionpair bonding, 113
Noradrenaline, 198
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role in Bruce effect, 201Norepinephrine (NE), see
NoradrenalineNotropis umbratili (redfin shiner fish)
interspecific egg dumper, 231Novomessor (ant)
recruitment, 140signals, 18
Nuptial dancepolychaete worms, 42
Odocoileus hemionus columbianus (black-tailed deer), see Deer
Odorant-binding proteins (OBP), 169,177, 178
Odour sieve, 167Oecophylla
Oecophylla longinoda (African weaverant)
alarm pheromone signal, 212, 214laying trail, 135territories, 88–89
Oecophylla smaragdinaterritories, 89
Oestrogen, see Hormones, oestrogenOestrus
Microtus ochrogaster (prairie vole), 198rats, 181, 191, 193, 196synchronisation
humans, 282–283, 285mice, 194rats, 196
Olfactionacross-fibre patterning, 171amygdala
interaction between main olfac-tory and vomeronasal organ(VNO) outputs, hamster, 184
behaviour of smelling (sniffs andflicks), 174
brainmacroglomerular complex (MGC)
not needed for response to allpheromones, 188
mapping activity, 172, 173,175–176, 185
moths and sex pheromones, 165,186–188
outputs modulated, 178, 182similarities in vertebrates and
insects, 165combinatorial coding, 170glomeruli, 170–174
integration of olfactory and visualinputs, 178
main olfactory systemfunctional overlap with
vomeronasal olfactory system(VNO), 180–86
mapping brain activity, fluorescing dyes, 172, 173functional magnetic resonance
imaging (fMRI)humans, 174, 175, 185
memories for odoursrenewal of olfactory sensory
neurons (OSNs), 173odotypes
hypothesis for coding features ofodour molecules, 170, 171
olfactory receptorstypes (and organisation)nematodes, insects, and
mammals, 172sensitivity
predator olfactory sensory neurons (OSNs) as sensitive asprey’s, 231
smell (olfaction) vs. taste (gustation), 164
specificityclean-up enzymes, 177odorant-binding proteins (OBPs),
177synaptic organization, 172
insect antennal lobe, 172mammalian olfactory bulb, 172
temporal coding in brains, 174fast oscillations of neural
activity, 174humans, 175fast oscillations, synchronisation,
174Olfactory bulb
ewe learns individual odour oflamb, 199, 200
Olfactory cuesrecognition
mates, species, kin, 198Olfactory organs
amphid sensillae (nematode),203–204
Argyropelecus hemigymnus (deep-seahatchet fish)
biggest nasal organ for body sizein vertebrates, 207
functional design, 166–168, 174lobsters
antennule sensory hairs, 174, 221sensitivity
dogs, 167moth antennae, 166
Olfactory receptor proteins (OR), 166evolution from G-protein-coupled
receptors, 176genes, 169–170
humans, 170Olfactory sensory neurons (OSNs),
165, 166renewal, 173similarity across animal
kingdom, 169stem cells, 169
Olfactory space, 169Oncorhynchus mykiss (rainbow trout)
taught to recognise predatorodour, 254
Ondata zibethicus (muskrats)monitoring, 257
Ophrys sphegodes (orchid)dupes solitary bee, 240, 242
Optical isomers, 305Optimal outbreeding
mate choice, 51Oreotragus oreotragus (klipspringer
antelope)scent marks eavesdropped by
ticks, 233Orientation behaviour, 206
active space, 209diffusion, 210–212
Argyropelecus hemigymnus (deep-seahatchet fish)
mate location, 207arresting, 207chemical plumes, 206combining information from
different senses, 223directly guided, 207idiothetic, 207, 208indirectly guided, 207, 208kinesis, 207, 208
klinokinesis, 208orthokinesis, 208
odour ‘landscape’, 206odour concentration gradient, 207odour stimulus
plumes, 210–227short range diffusion, 210–213, 214
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Orientation behaviour, (cont.)trails, 210, 213,214–218
precise stimuli, 207ranging, 209
strategies, 209sampling
sequential, 209simultaneous, 209
scalecommunication strategies,
212, 221self-steered, 207
counter-turning, 225taxis, 207, 208
anemotaxis, 222klinotaxis, 208rheotaxis, 222teleotaxis, 208tropotaxis, 208, 213, 222
teleology, 207threshold (K), 210turbulence
transmission in currents, 158virtual-reality chemical goggles, 223
Oryctolagus cuniculus (European rabbit)nipple pheromone, 181, 183
Ostariophysi, 158Ostrinia nubilalis (European corn borer
moth)pheromone signal genetics, 69
Ourebia ourebi (oribi), territories, 96Over-marking, 99–100
Castor canadensis (beaver), 96message centre, 99scent blending, 99scent masking, 99used in mate choice in house
mice? 94Oviposition-marking pheromones, see
Host-marking pheromonesOviposition pheromones
Simulidae (blackfly), 75Culex (mosquito), 75, 77Schistocerca gregaria (desert locust), 76
Ovis aries, see SheepOxytocin
peptide hormone, 199, 200
Pachycondyla (ants)Pachycondyla laevigata
mass recruitment, 134Pachycondyla obscuricornis
individual hunter, 134
Panorpa japonica (scorpion fly)symmetrical males more
attractive, 49Panulirus interruptus (California spiny
lobster)aggregation for defence, 75
Paraleptophlebia adoptiva (mayfly), 236Paraponera clavata (ant)
eavesdropped by parasitoid fly, 233Parasitoid insects
fliesApocephalus paraponerae
(phorid), 233Trichopoda pennipes (tachinid), 233
host suicide hypothesis, 152wasps
host marking, 84Telenomus euproctidis, 232Trichogramma pretiosum, 85
Parkinson’s disease, 297Paternal investment
Utetheisa ornatrix (tiger moth),47, 49, 50
Pectinophora gossypiella (pink bollwormmoth)
males, wider pheromone responsethan range females produce, 69
pest management, 258, 267Peptide pheromones, 15
aphrodisin, 182Litoria splendida (magnificent tree
frog), 21sodefrin, 15, 16specificity, 16
Perfumes, 273, 279Periglomerular cells, 172Periplaneta americana (American
cockroach)sex pheromone, 16
Periplanone-Bsex pheromone, Periplaneta
americana (Americancockroach), 16
Petaurus breviceps (sugar glider)scent marks affect sexual
activity, 196Petromyzon marinus (marine lamprey)
pest management, 262Pharmacophagy, 12Pheidole (ants)
Pheidole dentataenemy specification, 155
Pheidole pallidula
matching foraging effort to foodvalue, 139
Pheromone binding proteins(PBPs), 177
Pheromone identification, 25bioassays, 25–29
appropriate concentrations,28, 188
examples, 24, 25–29subtractive approach, 35Y-maze, 133
by comparative approachants, 32mice, 32
challenges, 23, 36collection of chemical signals, 29
aeration and cold trap, 24antelope scent marks, 29entrainment, Poropak-QTM,
Tenax™, cold trap, 31solid phase microextraction
(SPME), 31, 32, 33–34solvent washes or whole glands,
29, 31composite (combined) signals, 29electrophysiology
electroantennogram (EAG), 29, 31electroolfactogram (EOG), 29, 189electrovomerogram (EVG), 194single cell recording (SCR), 29
humansfunctional magnetic resonance
imaging (fMRI),174, 185organoleptic tests, 289‘scratch & sniff’ odorants, 297T-shirt sniff tests, 274twins studies, 292
ionised airtracking plumes, 221
model systems, 36, 203neurotransmitter dopamine as
plume tracer, 221new techniques
brain imaging, 36genetic analysis, 36genetic engineering, 35–36, 70genomics, 35
separating chemicalsfractionation, 25, 31gas liquid chromatography (GC or
GLC), 30, 32gas liquid chromatography–mass
spectroscopy (GCMS), 32
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high performance liquid chromatography (HPLC), 26
signal redundancydeer alarm signal, 28
Pheromones, 1advertising
Homarus americanus (lobster), 61aquatic habitats, 15assembly
guanine, tick, 80calling pattern, by mating system
and habitatstored-product insects, 82
caste differencesmandibular gland secretions,
114, 115changes with age
elephant males, 47humans, 275queen mandibular pheromone
(QPM) blend, 122convergent, 4–6cooperative breeding, 113–127
inhibition or suppression of subordinate reproduction, 126
coordinating externalfertilisation, 41
Nereis succinea (polychaeteworm), 42
Carassius auratus (goldfish), 9, 19costs of signalling, see Costscuticular hydrocarbons
dipteran sex pheromones, 70social insects, colony odours,
106, 108direct methods of transfer, 4
into bloodstream, 4to female, 4
dominance status, 44eavesdropped by visual cues, 232egg hatching pheromone (barnacle)
eicosanoids (PUFAs), 77evolution
alarm pheromones, 9clues from geographical
variation, 69, 232–234, 248eavesdropping, 8, 234effect of habitat, 82, 97–98enabled by olfactory receptor
proteins and combinatorialbrain circuits, 176
from existing chemical cues, 9,19, 42, 43, 189
honeybees, sociality, 114sensory drive, 6, 7
glandsmammals, summary, 10wasps, bees, ants, and termites,
summary, 11glycoproteins
barnacle settlement, 78salamander, 4
graded responsevole scent glands to
testosterone, 46hormone-based, 9host marking, 83–85insect
molecular weight related to function, 14
larval aggregation pheromonesbark beetles, 81
long distanceproduced by females, 41produced by males, 41
longevitymajor urinary proteins (MUPs), 193
major urinary proteins (MUPs)elephants, 195mice, 195
mammalsaphrodisin, 182detection by main olfactory
system, 181molecular weight related to
habitat, 13, 15marking of patches
bees, 85ladybirds, 85parasitoid wasps, 85
metabolic cost, lowAnthonomus grandis (boll weevil), 20Litoria splendida (magnificent tree
frog), 21molecular weight related to habitat
aquatic, 15terrestrial, 13, 15
multi-componentalarm pheromones in social
insects, 154moths, 17, 66provide specificity in insects and
vertebrates, 17recruitment, 141
multiple messages in odoursmammals, 102
social insects, 102nipple
Oryctolagus cuniculus (Europeanrabbit), 181, 183
oviposition, 75, 76, 77peptides
aphrodisin, 182larval settlement pheromone,
barnacle, 78Litoria splendida (magnificent tree
frog), 21marine invertebrates, 15pumping pheromone, crab, 77sodefrin, 15, 16
pheromone binding proteins (PBPs)polar molecules
anthopleurine, 15prostaglandins, 9pumping, 77queen egg marking, honeybee, 122queen mandibular pheromone
(QMP), 121, 204queen pheromones in social
insects, 118–123release as pulses rare, 221reproduction in social groups, 113sex ratio effects
Microcebus murinus (lesser mouselemur), 197
sourcesplants, 11, 12, 234symbiotic bacteria, 11, 111waste products, 80
stereoisomersimportance in invertebrates and
vertebrates, 17steroids, 9synergy in multi-component
pheromonesinsects, 35mammals, 35, 195
terrestrial habitats, 13–15transmission in currents
fish alarm pheromone, 158see also Aggregation pheromones;
Alarm pheromones; Antagonists; Glands; Honeybees; Host-markingpheromones; Pheromone identi-fication; Primer pheromones;Recruitment pheromones;Releaser pheromones; Sexpheromones; Signals; Territories
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386 INDEX
Philopatry, 125Phoxinus phoxinus (European minnow)
alarm pheromone, 158Pieris (cabbage butterfly)
host-marking pheromone, 84Pigs
minipig pets, 254see also Sus scrofa
Piriform cortex, 165Plankton,
commercial applications, disrupt-ing settlement, 266
settlementbarnacle, 26, 78Dendraster excentricus (sand
dollar), 80Hydroides (polychaete worm), 79
3D trailsTemora longicornis (copepod), 213
Plantsdeception by wild potatoes, 244bee pheromones from orchids, 12pyrrolizidine alkaloids (PAs), 12, 47tree defences against bark
beetles, 81see also Pheromones, sources, plants
Platysoma cylindrica, eavesdropping,234, 236
PlethodonPlethodon cinereus (red-backed
salamander)contest, 46feeding and breeding
territories, 46scent matching hypothesis and
territory, 95Plethodon jordani (salamander)
pheromone transfer by male, 4postcloacal gland, 95
Plotsus lineatus (catfish), 169Plumes
turbulence, visualisation, 219tracking plumes, 221
ionised air, 221neurotransmitter dopamine, 221
Podisus maculiventris (spined soldierbug)
eavesdropped by parasitoid, 233Pogonomyrmex badius (harvester ant)
alarm pheromone signal, 212Poison gland, ant, 130Polistes fuscatus (paper wasp)
kin recognition mechanisms, 107
Popilla japonica ( Japanese beetle), 17Postcloacal gland
salamander, 95Postpharyngeal gland
ants, 108host source of hydrocarbons for
parasite, 248Precopulatory mate guarding, see
Sperm competition and mateguarding
Predators and parasitoids, 2324-Pregnen-17�-,20�-diol-3-one
(17,20�-P), 9Pregnancy block, male pheromones
(Bruce effect), 195Premnas (anemone fish), 238Preorbital gland, see Antorbital glandPreputial gland
mice, 193Primates
Callithrix jacchus (common mar-moset)
subordinate female ovulationsuppressed, 197
Cerocebus albigena (mangabey), 271Leontopithecus rosalia (lion tamarin)
cooperative breeding not odourcontrolled, 127
Microcebus murinus (lesser mouselemur)
bias of offfspring sex ratios, 197female urine stimulates sper-
matogenesis and testosteronesecretion, 196
sexual activity of subordinatessuppressed, 196
micro-osmic, 271olfactory receptors, 169–170
genes, 170Saguinus fuscicollis (saddle-back
tamarin)multiple messages in odours, 102subordinate female ovulation
suppressed, 197scent glands and scent
marking, 272see also Humans
Primer pheromones, 18can be fast acting, 20Carassius auratus (goldfish), 19gregarising factor in egg foam
Schistocerca gregaria (desert locust), 76
juvenile hormone ( JH)social insects, 204ants, 204
maturation synchronySchistocerca gregaria (desert
locust), 76pest management, 266reproduction, 192
daylength and pheromones insheep, 192
similiarity of action in mammalsand insects, 20
social insectsqueen effects in ants, 204
termitescaste control, 119
Primiphales promelas (fathead minnow)alarm pheromone, 158
3-Propyl-1,2-dithiolane, 265Prostaglandins, 9
15-keto-prostaglandin-F-2�
(15-keto-PGF2�), 9fish peripheral response, 189
prostaglandin F-2� (PGF2�), 9Prostephanus truncatus (larger grain
borer beetle), 256males only signal until females
come, 82monitoring, 256
Proteles cristatus (aardwolf)‘dear-enemy’ individual
recognition, 113Protocerebrum, 165Proust, memory, 276Pseudaletia (moth), 5Pseudogates
termites, 119Pseudogenes, 170Puberty acceleration, see HormonesPUFAs (C20 polyunsaturated fatty
acids), 77Puntius (cyprinid fish), 189Pygidial gland, ant, 134Pyrrolizidine alkaloids (PAs), 12, 47
Queen mandibular pheromone (QMP),see Pheromones
Queen pheromones in social insects,see Pheromones
Racemate or racemic mixture, 306Ranging behaviour, 209Raphicerus melanotis (grysbok), 13
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Rattus norvegicus (Norway rat)follows trails from good food
sources, 137Recognition mechanisms, 103
direct familiarisation, 104indirect familiarisation (phenotypic
matching), 104Microtus ochrogaster (prairie vole),
197–198phenotypic matching hypothesis
field tests, 111recognition allele (‘green beard’
phenomenon), 104, 105fire ants, 105
sensitive period for learning, 105social insects
learning needed before discrimination, 103–109
usually by learning, 103–104Recruitment pheromones
alarmApis cerana japonica (Japanese
honeybee), 154, 156–157marking enemies for further
attack, 153, 154, 155ants
competition strategy, 140group recruitment, 134laying trail, 135mass communication, 129, 134scout ants, 134tandem running, 134trail specificity, 134worker sensitivity to trail
pheromone, 136aphids, social
soldiers attack enemy, 153, 153competition strategy, 140
ants, 140Trigona (stingless bee), 140
convergence in markingbehaviour, 135
coordinated attackVespa mandarinia japonica (giant
hornet), 154–155, 157expand colony diet, 136food distribution, recruitment
type, 136longevity related to food
supply, 137Malacosoma americanum (tent
caterpillar)laying trail, 135
matching foraging effort to foodvalue, 138
Rattus norvegicus (Norway rat)follow trails from good food
sources, 137scout ants, 134termites, 137
laying trail, 135trail specificity, 135
trail specificity, 140see also Alarm pheromones; Self-
organising systemsReinforcement, 65Releaser pheromones, 18
Carassius auratus (goldfish), 19mammal, 20nippleOryctolagus cuniculus (European rab-
bit), 181Reproductive character
displacement, 65Reproductive skew, 116, 117Response to pheromones, factors
affecting, 188, 191context
aphids tended by ants, 237honeybees, 188parasitoid wasps, 188
hormonescentral nervous system
effects, 189peripheral effects, 189
receiver’s characteristics, 188see also Learning; Development
Reticulitermies f lavipes (termite)recruitment pheromones, 140
Reynolds numbers (Re), 167copepod pheromone trails, 213
RhagolitisRhagolitis cerasi (cherry fruit fly)
pest management, 264Rhagolitis pomonella (apple
maggot fly)Crataegus mollis (hawthorn),
original host, 85evolution of host-marking
behaviour, 85experience of marking
pheromone needed forresponse, 190–191
host marking pheromone, 84Rhitropanopeus harrisii (mud crab)
pumping pheromone, 77
Ritualised signals, 3Ropalidia marginata (paper wasp)
non-kin may join colony, 108
Saguinus fuscicollis (saddle-backtamarin)
multiple messages in odours, 102subordinate female ovulation
suppressed, 197Salamanders and newts
courtship pheromone delivery patterns, 4
Cynops pyrrhogaster (red-belliednewt)
courtship, 16sodefrin, 16
Desmognathus ochrophaeusmale injects pheromone into
female, 4Plethodon cinereus (red-backed
salamander)postcloacal gland, 95scent matching, 95
Sanguinis fuscicollis (saddle-backtamarin)
multiple messages in odours, 102subordinate female ovulation
suppressed, 197Satellite males, see Alternative mating
stategiesSaturnia pavonia (emperor moth)
early demonstration ofpheromone, 23
Scaleboundary layer
effects on olfactory sensors, 174chemical plumes, 206communication strategies, 212design of olfactory organs, 166–168diffusable pheromones in water, 213Reynolds numbers (Re), 167, 213
Scent markingborder maintenance hypothesis, 90,
96–97Castor canadensis (beaver), 96ecological factors, patterns between
and within related species, 91males directly mark females
Dolichotis patagonum (mara), 100marking behaviour of house mice, 94non-territorial mammals, 100response by mice varies by
competitive ability, 94
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Scent marking (cont.)salamanders, nose-tapping, 95scent fence hypothesis, 91scent matching hypothesis, 90,
91–96mice, 93–94reducing cost of territorial
defence, 93salamanders, 95self-marking and presentation, 92testing the hypothesis, mice, 95
using vertebrate scent marks to census, 257
Schedorhinotermes (termites), 5Schedorhinotermes lamianus
foraging and soldiers/workers,138
pheromones guide food collection, 138
Schistocerca gregaria (desert locust)gregarising factor in egg foam, 76maturation synchrony, 76oviposition pheromone, 76pheromones in life cycle, 76
Scramble competition, 43–44Carassius auratus (goldfish), 43moth males responding to female
pheromones, 43selection for male moth sensitivity,
43, 166Thamnophis sirtalis parietalis
(red-sided garter snake), 43Sebaceous gland
birds, 21humans, 284signal life, 14
Sebum, 13Self-organising systems, 141
ant foraging, 141Eciton (army ants), 142matching resources to effort, 143modelling food distribution, 142stochastic effects, 143
termite nests, 143–145stigmergy, 143
‘Selfish herd’ theory, 159Semen
behavioural manipulation byDrosophila melanogaster, 63humans? 64Thamnophis sirtalis parietalis
(red-sided garter snake), 63–64Semibalanus balanoides (barnacle)
egg hatching pheromoneeicosanoids (PUFAs), 77
Semiochemicals, 1Sensitive periods, 201
ewe learning odour of her lamb,199, 200
hormonal state and learning, 199sex ratio effects
Microcebus murinus (lesser mouselemur), 197
termite caste development, 205Sensory drive, 6, 7Serricorone, 84Settlement-inducing protein complex
(SIPC), 78Settlement of marine vertebrates, 77–79Sex differences
in brainmacroglomerular complex (MGC)
in male moths, 190response to pheromones
Carassius auratus (goldfish), 189sensitivity to 3�-androstenol in
sows, 253see also Development
Sex pheromonesaphrodisin, 182Aplysia (sea-slug)
peptide sex pheromone, 15arachnids
Linyphia litogiosa (spider), 59crustaceans
Homarus americanus (lobster), 61, 62Temora longicornis (copepod), 213
exploitationcommercial, 253, 254natural, 231, 232–233
insectsCeratitis capitata (Mediterranean
fruit fly), 58, 59Lutzomyia longipalpis (sandfly), 58Nicrophorus (burying beetle), 59Periplaneta americana (American
cockroach), 16Xylocopa fimbriata (carpenter
bee), 57mammals, role of experience, 182sodefrin, Cynops pyrrhogaster (newt),
15, 16speciation, 64see also Development; Female
pheromones; Malepheromones
Sex ratioMicrocebus murinus (lesser mouse
lemur)bias of sex ratio in pregnancy,
197Sexual selection, 37
contests, same-sex, 38Homarus americanus (lobster), 44Nauphoeta cinerea (African cock-
roach), 40, 44female choice
elephants, 47Ephestia elutella (tobacco moth), 47Nauphoeta cinerea (African
cockroach), 40, 44male characters
coremata, 3, 10, 11, 47, 49mate choice, 37–38mate quality and courtship, 46mechanisms
direct benefit, 38, 39imprinting, 40indicator, 40runaway (‘Fisherian’, ‘sexy sons’),
38, 39, 59scramble competition, 43–44sexual competition, 38which sex should call? 40
‘Sexy faeces’ hypothesisPlethodon cinereus (red-backed
salamander), 46She-males, see Alternative mating
strategiesSheep (Ovis aries)
mate choice not universal, 56maternal behaviour, 199–201mother–infant recognition, 109reproduction, 192synchronised oestrus, 252
Signals, 1–2anonymous
social insects, 102butterflies
visual cues and pheromones, 14composite, 17–18
parallel sensory channels, 17with ultraviolet cues, 14
cooperative, 122design
aquatic habitats, 15contrasting modalities, 12habitat and daily activity, 13function and environment, 13
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lizard signals in contrasting habitats, 14
longevity, selection on, 13molecular weight and function,
13molecular weight and habitat,
13–15terrestrial habitats, 13
Dipsosaurus dorsalis (iguana)composite signal with ultraviolet
cue, 14dishonest, 121distinguishing chemical signals
from noise, 206honest, 3, 22, 44, 45
queen pheromoneshoneybees, 121, 121territorial scent marks, 89Utetheisa ornatrix (tiger moth),
47, 49, 49, 50low metabolic cost of pheromones
Anthonomus grandis (boll weevil), 20
Litoria splendida (magnificent treefrog), 21
mechanisms for specificitypeptides (amino acid sequence), 16unique molecules, 16multi-component, 17stereoisomerism, 17
modulationintensity, 17by sound, 18of visual threat by
pheromones, 18ON–OFF
clean-up enzymes, 177recruitment, 129
ants, 134redundancy, 17
deer alert signal, 28, 148ritualised, 3sexually selected, 37synergy, 35see also Alarm pheromones;
Pheromones, evolutionSimulidae (blackfly)
oviposition pheromone, 75, 263Singular breeders, 124Sitophilus (beetle), 83Smell, see OlfactionSnakes
Boiga irregularis (brown tree snake)
control, 262detecting trail direction, 217, 217Leptotyphlops dulcis (blind snake)
follows ant pheromone trails, 247tongue-flicking
trail following, 215Thamnophis sirtalis parietalis
(red-sided garter snake)overwintering aggregations, 80
Vipera berus (adder)Coolidge effect, 57
Sniffingbehaviour of smelling, 174
Social insectsalarm pheromones, 152–157ants
Atta texana, 136Oecophylla longinoda (African
weaver ant), 129, 212, 214Solenopsis invicta (fire ant), 123
aphidsalarm pheromones, 152–153Ceratovacuna lanigera, 153Colophina monstrifica, 153
caste differencespheromone secretions, 114reproductives and workers, 118response, 153
caterpillarsMalacosoma americanum (tent
caterpillar), 133colony odours
variation, 248guard bees
defense of colonies, 105discrimination of relatedness, 108recognition mechanisms, 105
Polistes fuscatus (paper wasps)kin recognition mechanisms, 107
Ropalidia marginatanon-kin may join colony, 108
parallels with social mammals, 127pheromones, 114primer pheromones
development of ant castes, 204queen mandibular pheromone
(QMP), 121action via juvenile hormone ( JH)
levels, 204queen pheromones, 118–123
control or cooperative signal, 118evolution, 122fire ants, 123
reproductive division of labour, 116termites
colony recognition, 109territories
Oecophylla longinoda (Africanweaver ant), 88–89
Oecophylla smaragdina (ant), 89Trigona (stingless bees)
odour beacons rather than trails,136
worker policing, 122see also Honeybees; Recruitment
pheromones; Self-organisingsystems
Social mammalsclan recognition, 111
Meles meles (European badger) , 111cooperative breeding, 124–127
evolution, 125inhibition or suppression of
subordinate reproduction, 126control by pheromones, 124, 126control not by pheromones, 124,
125, 127kin recognition
Castor canadensis (beaver), 111Spermophilus beldingi (Belding’s
ground squirrel), 110parallels with social insects, 112,
127primer pheromones and
reproduction, 123–124recruitment, 136
Heterocephalus glaber (naked molerats), 134
reproductive division of labour, 116shared colony odour, parallels
with social insects, 112Sodefrin, peptide pheromone, 15, 16Solanum berthaultii (wild potato),
255, 245Solenopsis (fire ants), 130, 137
Solenopsis invictaenemy specification, 155mass communication
recruitment, 134queen mutual inhibition, 123
Solid phase microextraction (SPME),31, 32
Speciation, 64–72allopatric, 64, 65Drosophila melanogaster
cuticular hydrocarbons, 70
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Speciation, (cont.)reproductive character
displacement, 65male pheromones, selection by
female to avoid hybrid matings, 70
signal shift, 68asymmetric tracking, 68
sympatric, 64, 65, 66Specificity of male moth responses to
pheromones, 68Specific-Mate Recognition System, 65Sperm competition and mate
guarding, 61–64cryptic female choice, 63
Tribolium castaneum (flour beetle),63
Utethiesa ornatrix (tiger moth), 63precopulatory mate guarding,
61–62consorting by male kangaroos, 62elephants, 62Homarus americanus (lobster), 61, 62
Spermophilus beldingi (Belding’sground squirrel)
kin recognition cues, 110Spindasis lohita (lycaenid butterfly),
238Squalene, 43‘Standing behaviour’, see LordosisStem cell, olfactory, 168–169Stereoisomers, 305Sternal gland
ant, 130Malacosoma americanum (tent
caterpillar), 130Polistes (paper wasp), 107termites, 130, 137
Steroids, 95�-cholestane 3,24-dione
Malacosoma trail pheromone, 130Stigmergy, see Self-organising systemsStimulo-deterrent diversionary
strategy (SDDS), 265Stochastic effects, 143Stored product insects
calling pattern, by mating systemand habitat, 82
Lasioderma serricorne (cigarette beetle), 83
Prostephanus truncatus (larger grainborer beetle) , 82, 256
Sitophilus (beetle), 83
Trogoderma granarium (Khapra beetle), 83
Stria terminalis, 183Subcaudal gland
Meles meles (European badger), 111Superparasitism, 83Sus scrofa (pig)
female response3�-androstenol, 20, 184, 2535�-androstenone, 184
lordosis by oestrous sow to malepheromones, 20, 184, 253
Swarming, see HoneybeesSweat glands
apocrine, 284eccrine, 284
Synergy, 35Synomones, 2
mutualism, 230
Tadpoles, kin, and alarmpheromones, 157
Talpa europaea (European mole), 91Taste
smell (olfaction) vs. taste(gustation), 164
taste buds, 164Taxis, 207, 208‘t-complex’ in mice, 56Telenomus euproctidis (parasitoid wasp)
eavesdrops host, 232Teleology, 207Temnochila chlorodia (beetle), 234Temora longicornis (copepod), 3D trails,
213Temporal gland
elephants, 46Termites
beetle mimics its termite host, 248caste-change pheromones, 119colony recognition odours, 109developmental pathways, 120pest management, 263primer pheromones, 119soldiers, control of numbers, 119
Termitophilous invertebrates, 245Territories, 87
costs (of signalling)Oreotragus oreotragus (klipspringer
antelope), 233dear-enemy phenomenon, 99economics of scent marking, 97
patterns, 97
variation by habitat inHyaenidae, 98
feeding and breedingPlethodon cinereus (red-backed
salamander), 46group territorial defence
Lemur catta (ring-tailed lemur)‘stink fights’, 90
‘owner advantage’Oecophylla longinoda (African
weaver ant), 88scent marking
border maintenance hypothesis,90, 96–97
composite signals withultraviolet cue, 99
lamp-post effect, 99social insects
Oecophylla longinoda (Africanweaver ant), 88–89
Testosterone, see Hormones,testosterone
Tetramorium caespitum (ant), foraging,143
Thamnophis sirtalis parietalis (gartersnake)
conflict between the sexessemen components, 63–64
detecting trail direction, 217, 217male pheromone, squalene, 43overwintering aggregations, 80she-males, 60
Thanasimus dubiuseavesdropping, 234
Thiazole, see Butyl dihydrothiazoleTicks
Amblyomma hebraeumpest management, 262
Dermacentor variabilispest management, 262
guanine, assembly pheromone, 80Ixodes neitzi responds to scent marks
of host antelope, 233Tools to study pheromones, see
Pheromone identificationTongue-flicking, snakes, 215Trail pheromones, see Recruitment
pheromonestrans (in chemical name), 305Tree defences, resins and toxic
chemicals, 81Tribolium castaneum (flour beetle), 82
cryptic female choice, 63
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Trichogramma pretiosum (parasitoidwasp), 85
Trichoplusia ni (cabbage looper moth)both sexes call, 41mutation in pheromone blend, 68male response requires hormone, 189
Trichopoda pennipes (tachinid)selection presure on host, 233
Trichopsenius frosti (beetle)matches its termite host, 248
(Z)-7-Tricosene, 61(E)-4-Tridecenyl acetate, 259Trigona (stingless bees), 5, 136, 140
Trigona angustularesists pheromone propaganda
by robber bee, 243Trigona subterranea
victim of pheromone propagandaby robber bee, 243
Trogoderma granarium (Khapra beetle),83
Trophallaxisoral, 119proctodeal, 119
Truffles (Tuber melanosporum)3�-androstenol, 253use of sows to find, 253human taste for, 288
Tuber melanosporum, see TrufflesTufted cells, 172
Ultraviolet (UV) cuesbirds eavesdrop vole scent marks
by UV, 232Dipsosaurus dorsalis (iguana),
composite signal with UV, 14Undercrowding
Allee effects, 75Urine
active only on contactMicrotus ochrogaster (prairie vole),
197–198dominant male effective, 192Homarus americanus (lobster)
contest, 44major urinary proteins (MUPs)
elephants, 195mice, 195
reflects hormonal and dominancestates in rodents, 46
Uropygidial gland, 21Utetheisa ornatrix (tiger moth)
female choice, 3, 63
honest signal, 47, 49, 49, 50hydroxydanaidal (HD) from diet, 47sexual selection, 39, 70
Vaginocervical stimulation duringbirth
hormonal releasers, 200Vandenbergh effect (puberty
acceleration), 192Vanillic acid, 260Varroa mite
pest management, 262response to honeybee larval
odours, 116Ventral amygdalofugal pathway, 183Verbenone, 235Veromessor (ant)
benzaldehyde as defensive compound, 5
Vertebratesandrogens
gland secretion rates, 46dual olfactory system, 178pheromones to attract mates, 41see also Amphibians; Antelopes;
Deer; Dogs; Fish; Mammals;Mice; Pheromones; Primates;Salamanders and newts;Social mammals; Wolves; othertaxonomic entries
Vespa mandarinia japonica (giant hornet), 154, 156–157
Vespula (wasps)Vespula germanica, 155
eavesdropping prey leks, 232, 231Vespula vulgaris, 155
Vipera berus (adder), Coolidge effect, 57Vomeronasal organ (VNO), 165,
178–185female VNO, urine contact, 198induction of oestrus
VNO needed, 181VNO not needed, 181
Vomeronasal amygdala, 165Vomeronasal olfactory system, 165
functional overlap with main olfactory system, 180–186
in humans? 294memory and pregnancy block
(Bruce effect) in mice, 201molecular architecture and
differences from main olfactory system (MOE),
179–180mapping to glomeruli, 180receptor proteins, 179receptor specificity, 179response to proteins, 180
some pheromones detected bymain olfactory system(MOE), 181
trail following by snakes, 215–217
Westermarck or kibbutz effecthuman mate choice, 280
Whitten effect (oestrus induction),192
WolvesCanis simensis (Ethiopian)
border marks in territory, 97Canis lupus (grey)
border marks in territory, 97control of reproduction, 124
Worker policing, 122
Xylocopa fimbriata (carpenter bee),leks, 57
Yponomeuta (small ermine moths)calling time, 66host plants, 66, 67pheromone blend, 66, 67
Zaspilothynnus trilobatus (thynninewasp)
duped by orchid, 241Zigzags
swimming or flying up plumes, 224Zootermopsis nevadensis (termite)
nymph laying trail, 135Zusammen, (Z) together, 308
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